DOCSLIB.ORG

Neuromechanism Study of Insect–Machine Interface: Flight Control by Neural Electrical Stimulation

Total Page：16

File Type：pdf, Size：1020Kb

Download full-text PDF Read full-text
Neuromechanism Study of Insect–Machine Interface: Flight Control by Neural Electrical Stimulation Neuromechanism Study of Insect–Machine Interface: Flight Control by Neural Electrical Stimulation Huixia Zhao1, Nenggan Zheng2*, Willi A. Ribi3, Huoqing Zheng1, Lei Xue2,4, Fan Gong2,4, Xiaoxiang Zheng2,4,5, Fuliang Hu1* 1 College of Animal Sciences, Zhejiang University, Hangzhou, China, 2 Qiushi Academy for Advanced Studies (QAAS), Zhejiang University, Hangzhou, China, 3 The Private University of Liechtenstein, Dorfstrasse 24, Triesen, Liechtenstein, 4 Department of Biomedical Engineering, Zhejiang University, Hangzhou, China, 5 Key Laboratory of Biomedical Engineering of Ministry of Education, Zhejiang University, Hangzhou, China Abstract The insect–machine interface (IMI) is a novel approach developed for man-made air vehicles, which directly controls insect flight by either neuromuscular or neural stimulation. In our previous study of IMI, we induced flight initiation and cessation reproducibly in restrained honeybees (Apis mellifera L.) via electrical stimulation of the bilateral optic lobes. To explore the neuromechanism underlying IMI, we applied electrical stimulation to seven subregions of the honeybee brain with the aid of a new method for localizing brain regions. Results showed that the success rate for initiating honeybee flight decreased in the order: a-lobe (or b-lobe), ellipsoid body, lobula, medulla and antennal lobe. Based on a comparison with other neurobiological studies in honeybees, we propose that there is a cluster of descending neurons in the honeybee brain that transmits neural excitation from stimulated brain areas to the thoracic ganglia, leading to flight behavior. This neural circuit may involve the higher-order integration center, the primary visual processing center and the suboesophageal ganglion, which is also associated with a possible learning and memory pathway. By pharmacologically manipulating the electrically stimulated honeybee brain, we have shown that octopamine, rather than dopamine, serotonin and acetylcholine, plays a part in the circuit underlying electrically elicited honeybee flight. Our study presents a new brain stimulation protocol for the honeybee–machine interface and has solved one of the questions with regard to understanding which functional divisions of the insect brain participate in flight control. It will support further studies to uncover the involved neurons inside specific brain areas and to test the hypothesized involvement of a visual learning and memory pathway in IMI flight control. Citation: Zhao H, Zheng N, Ribi WA, Zheng H, Xue L, et al. (2014) Neuromechanism Study of Insect–Machine Interface: Flight Control by Neural Electrical Stimulation. PLoS ONE 9(11): e113012. doi:10.1371/journal.pone.0113012 Editor: Paul Graham, University of Sussex, United Kingdom Received May 11, 2014; Accepted October 22, 2014; Published November 19, 2014 Copyright: ß 2014 Zhao et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Data Availability: The authors confirm that all data underlying the findings are fully available without restriction. All relevant data are within the paper and its Supporting Information files. Funding: The author FLH received the funding from the National Natural Science Foundation of China (No. 61031002), and the Modern Agroindustry Technology Research System from the Ministry of Agriculture of China (CARS-45). The author XXZ received the funding from the National Natural Science Foundation of China (No. 61003150), and the Fundamental Research Funds for the Central Universities. This work is partially supported by Zhejiang Provincial Natural Science Foundation of China (LZ14F020002). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. Competing Interests: The authors have declared that no competing interests exist. * Email: [email protected] (NZ); [email protected] (FH) Introduction behavior in locusts by a chronic electrical stimulation method on different brain sites [10]. And Blondeau (1981) had stimulated Insects, due to their impressive flight skills, are ranked among neurons in the lobula plate of free-moving and fixed Calliphora the best models for studying mechanisms of flight control, and for erythrocephala to evoke course control [11]. In these studies, use in the development of biomimetic micro air vehicles (MAVs) electrical stimulation was used as one of the tools of neuroethology [1]. As MAVs have become an increasingly hot topic of research, to investigate the relationship between animal behavior and the great advances have been made in the past few decades, but nervous system. Beyond that, electrical stimulation was also used significant challenges still remain with regards to payload mass, to artificially control the locomotion of an autonomous bio-robotic flight range, and speed [2,3]. A novel approach called the insect– system. Holzer and Shimoyama (1997) had induced the escape machine interface (IMI), which directly controls the flight behavior turn of cockroach via electrical stimulation to antennae, and built of insects by either neuromuscular or neural stimulation, has been an electronic backpack to control cockroach walking [12]. developed in recent years and promises to overcome some of the As to the IMI for MAVs and flight control, researchers challenges facing MAVs [4,5]. managed to elicit flight initiation and cessation in beetles (Cotinis The use of electrical stimulation to induce behavior in insects is texana and Mecynorhina ugandensis) by applying electrical not new. Singing behavior of the cricket and grasshopper had been stimulation through two electrodes implanted in the bilateral elicited by electrical stimulation of the brain [6,7] and descending optic lobes [13,14,15,16]. Their experiment was inspired by the fibres [8,9]. Rowell (1963) had produced various activities immediate flight cessation of untethered Mecynorhina ugandensis including antennal movements, locomotion, feeding, and sexual in response to abrupt darkening of the environment, which led the PLOS ONE | www.plosone.org 1 November 2014 | Volume 9 | Issue 11 | e113012 Flight Control in Honeybees researchers to hypothesize that the optic lobes might strongly Millonig’s buffer (pH 7.2) for 2–4 h before postfixation in a modulate flight initiation and cessation [5]. While alternating buffered 2% OsO4 solution for 1 h at room temperature. They positive and negative potential pulses at 100 Hz (20% duty cycle) were then dehydrated in an ethanol series (50%, 70%, 80%, 95%, initiated wing oscillations, a relatively long duration pulse caused 10 minutes each and 26100%, 15 minutes each), cleared twice in wing oscillations to cease [16]. Another study found that propylene oxide (15 and 20 minutes), and placed back into stimulating the antennal lobes with 20 Hz, 3.5 Vpp pulses initiated propylene oxide and Araldite mixture (1:1, 2–3 h) before being flight, while 50 Hz, 3.5 Vpp pulses stopped flight, in Manduca embedded in Araldite. Consecutive 1 mm thick sections were cut sexta [17]. In addition, ‘‘throttling’’ (frequency and stroke on a rotatory microtome using a diamond knife and stained with amplitude of wing oscillation) and turning modulation of insect toluidine blue. flight had been achieved by stimulating the optic lobes and basalar muscles [13,16], neck muscles [17] or ventral nerve cord Fixation and positioning system [18,19,20]. One of our previous studies developed a stimulation To assist with positioning the microelectrodes into specific brain protocol in restrained honeybees (Apis mellifera), using alternating subregions, we set up a fixation and positioning system by positive and negative electrical pulses (4 Vpp-40 Hz-40% duty assembling several pieces of equipment, including a honeybee cycle, t = 5 ms, m = 30) between two microwire electrodes clamping device (manufactured by our team), a digital stereotaxic implanted into the bilateral optic lobes to reproducibly generate apparatus (RWD Life Science Co., China), and a gimbaling flight initiation and cessation [21]. stereomicroscope with cold light illuminator (RWD Life Science It is difficult to elucidate the neuronal mechanisms of IMI from Co., China). The set-up is shown in Fig. 1. previous studies because identification of these mechanisms Honeybees were cold anesthetized before being fixed onto the requires electrophysiological or optical recordings of neuronal clamping device. A stereotyped procedure was applied to ensure activity in insects during stimulation [16]. We made some consistent head position of each fixed honeybee. When the rostral preliminary attempts to record neuronal activity, but unfortunately head was pressed down, the interspace between the retral head encountered problems due to interference from noise generated by and the clamp plates was filled up with beeswax–rosin mixture electrical stimulation of the brain (i.e. stimulus artifact) and robust (2:1). Meanwhile, special attention was paid to adjust the head avoidance behaviors of the insects (such as abdomen twitch and midline in accordance with a reference line on the clamp plates wing oscillations). We therefore posed an alternative question, (Fig. 1). After fixation, the honeybee underwent limb amputation namely how will stimulation of specific brain subregions (with the because legs gripping onto the clamp plates would hinder wing exception of the optic
Load more

Recommended publications
  • Coleoptera: Scarabaeidae: Cetoniinae): Larval Descriptions, Biological Notes and Phylogenetic Placement
    Eur. J. Entomol. 106: 95–106, 2009 http://www.eje.cz/scripts/viewabstract.php?abstract=1431 ISSN 1210-5759 (print), 1802-8829 (online) Afromontane Coelocorynus (Coleoptera: Scarabaeidae: Cetoniinae): Larval descriptions, biological notes and phylogenetic placement PETR ŠÍPEK1, BRUCE D. GILL2 and VASILY V. GREBENNIKOV 2 1Department of Zoology, Faculty of Science, Charles University in Prague, Viniþná 7, CZ-128 44 Praha 2, Czech Republic; e-mail: [email protected] 2Entomology Research Laboratory, Ottawa Plant and Seed Laboratories, Canadian Food Inspection Agency, K.W. Neatby Bldg., 960 Carling Avenue, Ottawa, Ontario K1A 0C6, Canada; e-mails: [email protected]; [email protected] Key words. Coleoptera, Scarabaeoidea, Cetoniinae, Valgini, Trichiini, Cryptodontina, Coelocorynus, larvae, morphology, phylogeny, Africa, Cameroon, Mt. Oku Abstract. This paper reports the collecting of adult beetles and third-instar larvae of Coelocorynus desfontainei Antoine, 1999 in Cameroon and provides new data on the biology of this high-altitude Afromontane genus. It also presents the first diagnosis of this genus based on larval characters and examination of its systematic position in a phylogenetic context using 78 parsimony informa- tive larval and adult characters. Based on the results of our analysis we (1) support the hypothesis that the tribe Trichiini is paraphy- letic with respect to both Valgini and the rest of the Cetoniinae, and (2) propose that the Trichiini subtribe Cryptodontina, represented by Coelocorynus, is a sister group of the Valgini: Valgina, represented by Valgus. The larvae-only analyses were about twofold better than the adults-only analyses in providing a phylogenetic resolution consistent with the larvae + adults analyses.
    [Show full text]
  • Coleoptera) with Corrections to Nomenclature and a Current Classification
    University of Nebraska - Lincoln DigitalCommons@University of Nebraska - Lincoln Papers in Entomology Museum, University of Nebraska State November 2006 A REVIEW OF THE FAMILY-GROUP NAMES FOR THE SUPERFAMILY SCARABAEOIDEA (COLEOPTERA) WITH CORRECTIONS TO NOMENCLATURE AND A CURRENT CLASSIFICATION Andrew B. T. Smith University of Nebraska - Lincoln, [email protected] Follow this and additional works at: https://digitalcommons.unl.edu/entomologypapers Part of the Entomology Commons Smith, Andrew B. T., "A REVIEW OF THE FAMILY-GROUP NAMES FOR THE SUPERFAMILY SCARABAEOIDEA (COLEOPTERA) WITH CORRECTIONS TO NOMENCLATURE AND A CURRENT CLASSIFICATION" (2006). Papers in Entomology. 122. https://digitalcommons.unl.edu/entomologypapers/122 This Article is brought to you for free and open access by the Museum, University of Nebraska State at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Papers in Entomology by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. Coleopterists Society Monograph Number 5:144–204. 2006. AREVIEW OF THE FAMILY-GROUP NAMES FOR THE SUPERFAMILY SCARABAEOIDEA (COLEOPTERA) WITH CORRECTIONS TO NOMENCLATURE AND A CURRENT CLASSIFICATION ANDREW B. T. SMITH Canadian Museum of Nature, P.O. Box 3443, Station D Ottawa, ON K1P 6P4, CANADA [email protected] Abstract For the first time, all family-group names in the superfamily Scarabaeoidea (Coleoptera) are evaluated using the International Code of Zoological Nomenclature to determine their availability and validity. A total of 383 family-group names were found to be available, and all are reviewed to scrutinize the correct spelling, author, date, nomenclatural availability and validity, and current classification status. Numerous corrections are given to various errors that are commonly perpetuated in the literature.
    [Show full text]
  • Coleoptera) with Corrections to Nomenclature and a Current Classification
    View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by Crossref University of Nebraska - Lincoln DigitalCommons@University of Nebraska - Lincoln Papers in Entomology Museum, University of Nebraska State November 2006 A REVIEW OF THE FAMILY-GROUP NAMES FOR THE SUPERFAMILY SCARABAEOIDEA (COLEOPTERA) WITH CORRECTIONS TO NOMENCLATURE AND A CURRENT CLASSIFICATION Andrew B. T. Smith University of Nebraska - Lincoln, [email protected] Follow this and additional works at: https://digitalcommons.unl.edu/entomologypapers Part of the Entomology Commons Smith, Andrew B. T., "A REVIEW OF THE FAMILY-GROUP NAMES FOR THE SUPERFAMILY SCARABAEOIDEA (COLEOPTERA) WITH CORRECTIONS TO NOMENCLATURE AND A CURRENT CLASSIFICATION" (2006). Papers in Entomology. 122. https://digitalcommons.unl.edu/entomologypapers/122 This Article is brought to you for free and open access by the Museum, University of Nebraska State at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Papers in Entomology by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. Coleopterists Society Monograph Number 5:144–204. 2006. AREVIEW OF THE FAMILY-GROUP NAMES FOR THE SUPERFAMILY SCARABAEOIDEA (COLEOPTERA) WITH CORRECTIONS TO NOMENCLATURE AND A CURRENT CLASSIFICATION ANDREW B. T. SMITH Canadian Museum of Nature, P.O. Box 3443, Station D Ottawa, ON K1P 6P4, CANADA [email protected] Abstract For the first time, all family-group names in the superfamily Scarabaeoidea (Coleoptera) are evaluated using the International Code of Zoological Nomenclature to determine their availability and validity. A total of 383 family-group names were found to be available, and all are reviewed to scrutinize the correct spelling, author, date, nomenclatural availability and validity, and current classification status.
    [Show full text]
  • Recent Developments in the Remote Radio Control of Insect Flight
    FOCUSED REVIEW published: 08 December 2010 doi: 10.3389/fnins.2010.00199 Recent developments in the remote radio control of insect flight Hirotaka Sato* and Michel M. Maharbiz* Department of Electrical Engineering and Computer Science, University of California at Berkeley, Berkeley, CA, USA The continuing miniaturization of digital circuits and the development of low power radio systems coupled with continuing studies into the neurophysiology and dynamics of insect flight are enabling a new class of implantable interfaces capable of controlling insects in free flight for extended periods. We provide context for these developments, review the state-of-the-art and discuss future directions in this field. Keywords: insect flight, micro air vehicle (MAV), telemetry, cyborg insect, brain machine interface (BMI) Edited by: Rui M. Costa, Instituto Gulbenkian de Ciência, Portugal INTRODUCTION and Boettiger, 1965; Leston et al., 1965; Ellington, Reviewed by: Ty Hedrick, The University of North Insects exhibit some amazing modes of locomo- 1984; Tu and Dickinson, 1994, 1996; Ellington Carolina at Chapel Hill, USA tion. They float, swim, crawl, jump, run (even et al., 1996; Dickinson and Tu, 1997; Dickinson M. Eugenia Chiappe, Howard Hughes bipedally), and fly. They do this with such effi- et al., 1999, 2000; Josephson et al., 2000a,b) engi- Medical Institute, USA ciency, sophistication, and elegance of function neers are starting to build tiny, sub-gram scale, *Correspondence: across a vast size range (the smallest insect is flying robots. 1/5 of a millimeter across, the largest insects can Building very small flyers which can fly well weigh 100 g and be over 20 cm long) that they in real environments is a difficult task, however, have mastered locomotion on almost every ter- hampered not just by knowledge of insect flight, rain on Earth.
    [Show full text]
  • Authorsindex
    AUTHORSINDEX ARROW, G. J. 153. HUXLEY, J. 103, 125, 153. BAKER, C. S. 59. ILLIGER, J. C. 27. BATES, H. W. 76, 153. JUDULIEN, F. 55, 154. BATESON, W. 60, 102, 153. LAMEERE, A. 102, 133, 136, 139, 154. BEEBE, W. 61 et seq., 132, 153. LOCKET, G. H. 126, 154. BEESON, C. F. C. 37, 153. MABTERLINCK, M. 29. BENNETT, A. L. 63, 153. MAIN, H. 11, 43, 154. BLAIR, K. G. 1. MANEE, A. H. 45, 57, 59, 154. BRINDLEY, H. H. 60, 102, 153. MARSHALL, SIR G. A. K. 1, 75. BRISTOWE, W. S. 126, 153. MORLEY, B. D. W. 123, 154. CHAMPY, C. 138, 153. MULSANT, E. 27, 154. CHAPMAN, T. A. 34 et seq., 38, 153. OHAUS, F. 29 et seq., 54, 56 et seq., CORBIN, G. B. 42, 153. 69, 98, 154. CORNELIUS, C. 141. PEARSE, A. S. 31, 154. CUNNINGHAM, J. T. 124, 127, 153. PECKHAM, G. & E. 125, 154. DAGUERRE, J. B. 55, 153. PERCIVAL, R. B. 112. DARWIN, C. 2, 3, 24, 65, 79, 103, 108, PERRIS, E. 79. 116 et seq., 132, 139, 153. POCOCK, R. I. 1. DOANE, R. W. 58, 153. REICHENAU, W. VON. 118, 154. ELTRINGHAM, H. 119. RODWAY, F. A. 48. EMDEN,F.VAN. 1. SAUNDERS, C. J. 48, 154. FASRE, J. H. 8, 11, 27, 28, 43, 49 et SCHREINER, J. 39, 154. seq., 74, 153. SLOANE, H. 53. FRASER, F. C. 31. STROHMEYER, H. 13, 154. GEDDES, SIR P. 118, 139, 154. SYMS, E. 74. HEYMONS, R. 31, 153. THOMSON, SIR J. A. 118, 139, 154.
    [Show full text]
  • A New Key to the Suprageneric Taxa in the Beetle Family Cetoniidae, with Annotated Lists of the Known Genera
    A NEW KEY TO THE SUPRAGENERIC TAXA IN THE BEETLE FAMILY CETONIIDAE, WITH ANNOTATED LISTS OF THE KNOWN GENERA by J. KRIKKEN J. Krikken: A new key to the suprageneric taxa in the beetle family Cetoniidae, with annota• ted lists of the known genera. Zool. Verh. Leiden 210, 5-ix-1984: 1-75, figs. 1-86, 1 table. — ISSN 0024-1652. Key-words: Coleoptera, Scarabaeoidea, Cetoniidae; key, family-group taxa; checklist, genera. A revised key to the subfamilies, tribes and subtribes of the Cetoniidae (Coleoptera: Scara• baeoidea) is presented. Comments are given on the classification implied by the key, which inclu• des several redefinitions. All the known genera (ca. 510) are listed under their tribe or subtribe, the position of several being different from that in Schenkling's catalogues (1921, 1922). Referen• ces to recent synoptic literature are given. Eighteen new subtribes are proposed, the majority in the tribe Cremastocheilini. One new tribe is proposed, the Platygeniini. Attention is drawn to various new and neglected synonymies, original spellings, and other nomenclatural matters regar• ding family-group and genus-group names. The position of many genera is briefly discussed. Some striking features of the biogeography of the Cetoniidae are discussed, such as the high ge• neric endemicity for the major biogeographical regions (for the Madagascan, Afrotropical and Australasian regions ca. 90% or more). J. Krikken, Rijksmuseum van Natuurlijke Historie, P.O. Box 9517, 2300 RA Leiden, the Ne• therlands. CONTENTS 1. INTRODUCTION 4 Purpose, limitations, cetoniid natural history 4 General taxonomic and nomenclatural comments 7 Technical explanation 9 Biogeographical summary 11 2.
    [Show full text]
  • Radio-Controlled Cyborg Beetles: a Radio-Frequency System for Insect Neural Flight Control H
    RADIO-CONTROLLED CYBORG BEETLES: A RADIO-FREQUENCY SYSTEM FOR INSECT NEURAL FLIGHT CONTROL H. Sato1, Y. Peeri1, E. Baghoomian1, C.W. Berry2, M.M. Maharbiz1,2 1Electrical Engineering and Computer Science, University of California, Berkeley, CA, USA 2Electrical Engineering and Computer Science, University of Michigan, Ann Arbor, MI, USA ABSTRACT 1. INTRODUCTION We present the first report of radio control of a cyborg Micro air vehicles (MAV’s) which can navigate into beetle in free-flight. The microsystem (Figs. 1, 2) consisted locations not easily accessible to humans have been the of a radio-frequency receiver assembly, a micro battery and subject of much recent research [1]. However, man-made a live giant flower beetle platform (Mecynorhina MAV’s are still limited in size, payload, distance and polyphemus or Mecynorhina torquata). The assembly had performance. In contrast, many insects have as-yet six electrode stimulators implanted into the left and right unmatched flight performance and increasingly understood optic lobes, brain, posterior pronotum (counter electrode), muscular and nervous systems [2]. Additionally, some right and left basalar flight muscles. Initiation and insects undergo complete metamorphosis (i.e. form pupae) cessation of flight were accomplished by optic lobe and are amenable to implantation and internal stimulation while muscular stimulation of either right or manipulation during pupation. In light of this, there have left basalar flight muscles (referenced to the posterior been recent efforts by several groups to implant pronotum electrode) elicited left or right turns, respectively. microsystems into insects to control their flight [3-8]. Flight commands were wirelessly transferred to the beetle-mounted system (running BeetleBrain v1.0 code) via an RF transmitter operated by a laptop running custom software (BeetleCo mmander v1.0) through a USB/Serial interface.
    [Show full text]
  • Immature Stages of Giants: Morphology And
    A peer-reviewed open-access journal ZooKeys 619: 25–44Immature (2016) stages of giants: morphology and growth characteristics of Goliathus... 25 doi: 10.3897/zookeys.619.8145 RESEARCH ARTICLE http://zookeys.pensoft.net Launched to accelerate biodiversity research Immature stages of giants: morphology and growth characteristics of Goliathus Lamarck, 1801 larvae indicate a predatory way of life (Coleoptera, Scarabaeidae, Cetoniinae) Tomáš Vendl1, Petr Šípek1 1 Department of Zoology, Faculty of Science, Charles University, Viničná 7, CZ-128 44 Praha 2, Czech Republic Corresponding author: Petr Šípek ([email protected]) Academic editor: F. Krell | Received 16 February 2016 | Accepted 26 August 2016 | Published 27 September 2016 http://zoobank.org/0AB2CB66-7539-4A5B-AC5C-8C37DDF7CDC2 Citation: Vendl T, Šípek P (2016) Immature stages of giants: morphology and growth characteristics of Goliathus Lamarck, 1801 larvae indicate a predatory way of life (Coleoptera, Scarabaeidae, Cetoniinae). ZooKeys 619: 25–44. doi: 10.3897/zookeys.619.8145 Abstract The third larval instar of Goliathus goliatus (Drury, 1770), G. orientalis Moser, 1909 and G. albosignatus Boheman, 1857 are described and illustrated for the first time and compared with the immature stages of other Cetoniinae. Larval development of G. goliatus is investigated under laboratory conditions, with particular emphasis on food requirements. These results support the obligatory requirement of proteins in the larval diet. The association between larval morphological traits (e. g., the shape of the mandibles and pretarsus, presence of well-developed stemmata) and larval biology is discussed. Based on observations and the data from captive breeds it is concluded that a possible shift from pure saprophagy to an obligatory predaceous way of larval life occurred within the larvae of this genus, which may explain why these beetles achieve such an enormous size.
    [Show full text]
  • Hidden Complexity in the Ontogeny of Sexual Size Dimorphism in Male-Larger Beetles
    www.nature.com/scientificreports OPEN Hidden complexity in the ontogeny of sexual size dimorphism in male- larger beetles Received: 27 October 2017 Tomáš Vendl1,3, Petr Šípek1, Ondřej Kouklík1 & Lukáš Kratochvíl2 Accepted: 22 March 2018 Sexual size dimorphism (SSD) is widespread among animals, but its developmental mechanisms are not Published: xx xx xxxx fully undestood. We investigated the proximate causes of SSD in three male-larger and one monomorphic scarab beetles using detailed monitoring of growth in individual instars. Apart from the fnding that SSD in all three male-larger species started to develop already in the frst larval instar, we generally found a high variability in SSD formation among the species as well as among instars. Overall, sexual diferences in developmental time, average growth rate, as well as in the shape of the growth trajectory seem to be the mechanisms responsible for SSD ontogeny in scarab beetles. In the third instar, when the larvae attain most of their mass, the males had a similar or even lower instantaneous growth rate than females and SSD largely developed as a consequence of a longer period of rapid growth in males even in cases when the sexes did not difer in the total duration of this instar. Our results demonstrate that a detailed approach, examining not only the average growth rate and developmental time, but also the shape of the growth trajectory, is necessary to elucidate the complex development of SSD. Female-biased sexual size dimorphism (SSD), i.e. situation when females are the larger sex, is prevalent in poikilothermic vertebrates1,2 and most arthropods, including insects3,4.
    [Show full text]
  • Nhbs Monthly Catalogue New and Forthcoming Titles Issue: 2010 Complete January 2011 [email protected] +44 (0)1803 865913
    nhbs monthly catalogue new and forthcoming titles Issue: 2010 complete January 2011 www.nhbs.com [email protected] +44 (0)1803 865913 Zoology: The NHBS Monthly Catalogue in a complete yearly edition Mammals Birds Reptiles & Amphibians Welcome to the Complete 2009 edition of the NHBS Monthly Catalogue, the ultimate Fishes buyer's guide to new and forthcoming titles in natural history, conservation and the Invertebrates environment. With 300-400 new titles sourced every month from publishers and Palaeontology research organisations around the world, the catalogue provides key bibliographic data Marine & Freshwater Biology plus convenient hyperlinks to more complete information and nhbs.com online General Natural History shopping - an invaluable resource. Each month's catalogue is sent out as an HTML Regional & Travel email to registered subscribers (a plain text version is available on request). It is also Botany & Plant Science available online, and offered as a PDF download. Animal & General Biology Please see our info page for more details, also our standard terms and conditions. Evolutionary Biology Prices are correct at the time of publication, please check www.nhbs.com for the latest Ecology prices. Habitats & Ecosystems Conservation & Biodiversity Environmental Science Physical Sciences Sustainable Development Data Analysis Reference Mammals Go to subject web page A l'Affut des Loutres 120 pages | 98 photos | BIOTOPE Stephane Raimond Parthenope Contains the author's colour photos of otters in the Millevaches plateau, part of the Limousin Pbk | 2009 | 2917848049 | #186086A | region of France. .... £27.99 Add to basket Animal Reintroductions 291 pages | Figs, tabs | CUP The Arabian Oryx in Oman Pbk | 2010 | 0521131677 | #184338A | Mark R Stanley Price £19.99 Add to basket When this book was first published in 1989, attempts to reintroduce plants or animals into their native habitat were still relatively rare, and those that were adequately ...
    [Show full text]
  • A Survey of the Rhinoceros Beetle and Stag Beetle Market in Japan
    Published by TRAFFIC East Asia-Japan,Tokyo,Japan 出版元：トラフィックイーストアジアジャパン、東京 ○C 2003 TRAFFIC East Asia-Japan ○C 2003トラフィックイーストアジアジャパン All rights reserved. このレポートの著作権はすべてトラフィックイーストアジ アジャパンに属します。 All material appearing in this publication is copy- 本報告書の無断転載はお断り致します。 righted and may be reproduced with permission. 転載ご希望の際はトラフィックイーストアジアジャパンに Any reproduction in full or in part of this publication ご一報ください。 must credit TRAFFIC East Asia-Japan as the copy- right owner. The views of the author expressed in this publication このレポートの著者の意見は、必ずしもトラフィックイー do not necessarily reflect those of the TRAFFIC East ストアジア、WWFまたはIUCNの意見を反映しているとは限 Asia, WWF or IUCN. りません。 The designations of geographical entities in this pub- このレポートの中での地理的名称、および資料の表記は、 lication, and the presentation of the material, do not いかなる国、領土、地域、当局の法律の現状、もしくは境 imply the expression of any opinion whatsoever on 界、国境の設定に関するトラフィックまたは、その支援機 the part of TRAFFIC or its supporting organizations 関の意見を反映するものではありません。 concerning the legal status of any country, territory, or area, or of its authorities, or concerning the delimi- tation of its frontiers or boundaries. The TRAFFIC symbol copyright and Registered トラフィックのシンボルの著作権、登録商標の所有権は Trademark ownership is held by WWF. TRAFFIC is WWFに属します。トラフィックはWWFとIUCNの共同プロ a joint programme of WWF and IUCN. グラムです。 Suggested citation: Kameoka, S. and Kiyono, H. 引用例: 亀岡晶子・清野比咲子(2003). カブトムシとクワガタ (2003).A Survey of the Rhinoceros Beetle and Stag ムシの市場調査 トラフィック イーストアジア ジャパン Beetle Market in Japan. TRAFFIC East Asia- Japan. ISBN:4-902548-00-3 文書コード：4-902548-00-3 Front cover:
    [Show full text]
  • Insects in Food and Feed Systems in Sub-Saharan Africa: the Untapped Potentials
    International Journal of Tropical Insect Science https://doi.org/10.1007/s42690-020-00305-6 MINI-REVIEW Insects in food and feed systems in sub-Saharan Africa: the untapped potentials Samuel A. Babarinde1 & Brighton M. Mvumi2 & Grace O. Babarinde3 & Faith A. Manditsera4 & Taiwo O. Akande5 & Adebusola A. Adepoju6 Received: 31 May 2020 /Accepted: 28 September 2020 # African Association of Insect Scientists 2020 Abstract Insects as food and feed have the potential to alleviate food, feed and nutrition insecurity in sub-Saharan Africa (SSA) against a backdrop of climate change. Such use has gained unprecedented attention in the past decade and the trend will probably continue due to the species diversity, new discoveries in the nutritional, neutraceutical and medicinal potentials of edible insect species. In order to meet the increasing demand for insects as food and feed, insect farming should complement sustainable wild insect harvesting. The ecological impact of insect farming, economics, species biological and processing aspects deserve empirical investigation. This is crucial in order to effectively guide potential insect producers and processors. Besides the use of insects in folk medicine, several industrial products including polyunsat- urated and monosaturated fatty acids, peptides, enzymes, and antimicrobial compounds can be obtained from edible insects. With the teaming world population, value addition via product fortification is a practical strategy to enhance the acceptance of edible insects for human food and nutrition security. The future of insects as food and feed will witness the development of international trade and SSA governments should be ready to comply with product standardization and legislation requirements to penetrate external markets.
    [Show full text]